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Flying insects have elastic materials within their exoskeletons that could reduce the energetic cost of flight if their wingbeat frequency is matched to a mechanical resonance frequency. Flapping at resonance may be essential across flying insects because of the power demands of small-scale flapping flight. However, building up large-amplitude resonant wingbeats over many wingstrokes may be detrimental for control if the total mechanical energy in the spring-wing system exceeds the per-cycle work capacity of the flight musculature. While the mechanics of the insect flight apparatus can behave as a resonant system, the question of whether insects flap their wings at their resonant frequency remains unanswered. Using previous measurements of body stiffness in the hawkmoth, Manduca sexta , we develop a mechanical model of spring-wing resonance with aerodynamic damping and characterize the hawkmoth's resonant frequency. We find that the hawkmoth's wingbeat frequency is approximately 80% above resonance and remains so when accounting for uncertainty in model parameters. In this regime, hawkmoths may still benefit from elastic energy exchange while enabling control of aerodynamic forces via frequency modulation. We conclude that, while insects use resonant mechanics, tuning wingbeats to a simple resonance peak is not a necessary feature for all centimetre-scale flapping flyers.
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Rapid frequency modulation in a resonant system: aerial perturbation recovery in hawkmoths
Centimetre-scale fliers must contend with the high power requirements of flapping flight. Insects have elastic elements in their thoraxes which may reduce the inertial costs of their flapping wings. Matching wingbeat frequency to a mechanical resonance can be energetically favourable, but also poses control challenges. Many insects use frequency modulation on long timescales, but wingstroke-to-wingstroke modulation of wingbeat frequencies in a resonant spring-wing system is potentially costly because muscles must work against the elastic flight system. Nonetheless, rapid frequency and amplitude modulation may be a useful control modality. The hawkmoth Manduca sexta has an elastic thorax capable of storing and returning significant energy. However, its nervous system also has the potential to modulate the driving frequency of flapping because its flight muscles are synchronous. We tested whether hovering hawkmoths rapidly alter frequency during perturbations with vortex rings. We observed both frequency modulation (32% around mean) and amplitude modulation (37%) occurring over several wingstrokes. Instantaneous phase analysis of wing kinematics revealed that more than 85% of perturbation responses required active changes in neurogenic driving frequency. Unlike their robotic counterparts that abdicate frequency modulation for energy efficiency, synchronous insects use wingstroke-to-wingstroke frequency modulation despite the power demands required for deviating from resonance.
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- Award ID(s):
- 1806833
- PAR ID:
- 10282870
- Date Published:
- Journal Name:
- Proceedings of the Royal Society B: Biological Sciences
- Volume:
- 288
- Issue:
- 1951
- ISSN:
- 0962-8452
- Page Range / eLocation ID:
- 20210352
- Format(s):
- Medium: X
- Sponsoring Org:
- National Science Foundation
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Accepted Manuscript1.0
- Journal Article:
- https://doi.org/10.1098/rspb.2021.0352
